Saura Hereditas (2020) 157:48 https://doi.org/10.1186/s41065-020-00161-x

REVIEW Open Access Race Anssi Saura

Abstract Background: The founders of Hereditas envisioned that race biology would be a major subject that had social applications with utmost importance in the near future. Anthropometrics was in this context understood to be the pure and the applied science. had a long tradition in physical . Herman Lundborg, member of the advisory board of Hereditas, united the anthropometric and eugenic approaches in a synthesis. He was the first head of the Institute for Race Biology in Sweden. The contents of Hereditas reflect the development of race biology in the Nordic countries. Conclusions: The initial enthusiasm for applied race biology did not last long. In the 1920’s Hereditas carried papers on both physical and eugenics. Most paper dealt, however, with human without eugenic content. Two papers, published in 1921 and 1939 show how the intellectual climate had changed from positive to negative. Finally only human genetics prevailed as the legitimate study of the human race or humankind. A belated defense of eugenics published in 1951 did not help; geneticists had abandoned anthropometrics for good around the year 1940 and eugenics about a decade later. In spite of that, eugenic legislation was amended astonishingly late, in the 1970’s. The development was essentially similar in all Nordic countries. Keywords: anthropometrics, eugenics, sterilization, Hereditas

Background Race biology stands for biological anthropology with A race is a population distinguishable from other popu- eugenics as an applied approach. In the Nordic countries lations by hereditary phenotypic characters. Members of the two make up a temporal sequence so that anthropo- a human race may freely produce offspring with repre- logical measurements came first, followed with the social sentatives of other races. Even if race has been socially aspect. constructed [1], it has been an immensely powerful con- Linnaeus had in Systema naturae divided humanity struction that has shaped the lives of millions. Restric- into four differentiated groups using geographic but tions based on geography, skin color, language, culture otherwise arbitrary criteria. A systematic research on and religion have been imposed to maintain groups of race was begun in Sweden in the 1840’s. The father of people as endogamous units. The Indian caste system anthropometrics is Anders Rezius (1796–1869), who may have been the most extreme effort of social control took measures on Sámi (Lapps) [3, 4] . He introduced of human reproduction [2]. Segregation in the American , the study of skull dimensions, and coined south, apartheid in South Africa, aristocratic societies terms such as dolichocephalic and brachycephalic, long- and prejudices represent less brazen attempts of social and short-skulled, respectively. His work led to a revi- control. The Universal Declaration of Human Rights was sion of race classification in Europe. Gustaf Retzius drafted to abolish the above practices. (1842–1919) followed his father as an anthropologist. He was interested in the three races of Sweden: Nordic, Finnish and Sámi (Lapps). The Nordics were in general Correspondence: [email protected] dolichocephalic, the Sámi brachycephalic and the Finns Department of Molecular Biology, Umeå University, SE-901 87 Umeå, Sweden

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intermediate between the two. Typical Sámi were seen a collection of photos and paintings showing Swedes to- to have a short stature and dark hair [5], while the gether with Sámi, deviant types and criminals. The ex- Swedes and Finns were tall and had fair hair. Retzius jr. planatory text included the elements of genetics. Starting was, in particular, keen to measure Finnish skulls, [6]. from and , the exhibition toured sev- The giant volume titled Anthropologia Suecica [7] sums eral towns in 1919. It was an all-time success that up the accumulated results of 60 years. In retrospect, it attracted more than 50% of the adult population in many is difficult to make out why this immoderate compilation cities [14]. That is no wonder: Anders Zorn specialized in of data was collected. It was certainly used to promote painting nude blond maidens displaying their charms in ideas of the pre-eminence of the “superior dolicocephalic lush lakeside etc. settings. race” e.g. [8]. Given this alleged superiority of the , posi- Hereditas tive measures had to be taken not only to preserve it at Herman Nilsson-Ehle, the plant breeder of international the degree of purity then attained and, if feasible, to im- fame, was the driving force in the founding of the Men- prove the positive and desirable and weed out the nega- delian Society in Lund in 1910 and, ten years later, in tive characteristics. Herman Lundborg extended the setting up its journal Hereditas. He was involved in vir- work of Retzius and Fürst [7] into a medical dimension. tually everything that was done to further the progress He [9] defended in 1903 a doctoral dissertation on myo- of genetics in Sweden between the years 1910 and 1935 clonus epilepsy or Unverricht-Lundborg disease. Lund- [15–17]. To secure a financial base for a genetics journal borg used pedigrees extracted from church records to the Mendelian Society sent an appeal to Swedish patrons find out the lines of descent and the transmission of the of science and culture. The tone of the appeal, signed disease. His thesis represented a new and useful ap- by Herman Nilsson-Ehle, president, and Carl August proach in the study of heritable diseases. Lundborg was Hallqvist, secretary general, was patriotic. They stressed rewarded with accolades including the prize of the the significance of genetics in plant breeding and applied Swedish Medical Society. The recognition encouraged race biology; the plant breeding institutions of Svalöf and him to extend the study beyond strict human genetics. Weibullsholm were taken up by name. Race biology has In a follow-up study comprising an extended family with social applications that will be of utmost importance in 2232 members Lundborg [10] not only demonstrated the near future. If diseases and traits are found to have a the inheritance of the epilepsy but, in addition, contro- genetic basis, then the society should respond through ap- versial evidence for degeneration that expressed itself as propriate measures. Financial utility is less important than alcoholism and debauchery. Cousin marriages had exac- the control of asocial behavior etc. Evidently a critical and erbated the misfortune. Lundborg recommended that strictly scientific treatment is needed here [16, 17]. eugenic measures should be applied to cure the afflicted Herman Lundborg was elected member of the advisory family. The result was a giant volume (35.5 × 28× 5.5 cm, board of Hereditas from almost the very beginning. Heredi- 740 pages). In addition to his own data, he described the tas was envisioned to be a publishing forum for Nordic ge- racial characters of the Swedish people following Retzius neticists so that Swedes, and authors of PhD theses in and Fürst [10]. particular, were given priority, but if space allowed, contri- The Swedish Society for Race Biology (Svenska Sällska- butions from other Nordic countries were included. pet för Rashygien) was established in the autumn of 1909 in Stockholm. Rashygien,“race hygiene”, is a German and Applied race biology Nordic synonym for eugenics, albeit with some anti- Theodore M Porter [18] has pointed out that physicians Semitic overtones [11]. The society had a lofty aim: “to and state officials tasked with overseeing insane asylums support efforts to improve the health of the body and throughout the nineteenth century attempted to under- mind in the future generations”. The evolutionary biolo- stand the nature and origins of madness. They were the gist Wilhelm Leche was the chairman and the Nobel Prize precursors of pedigree research and eugenics. winner (chemistry) Svante Arrhenius the most prominent In the first half of the twentieth century eugenics was member of the executive board. Herman Lundborg a legitimate branch of applied genetics. Negative eugenics emerged as an active advocate for race biology. Together strove to lower the frequency of detrimental traits. Indi- with the zoologist Nils von Hofsten, the psychiatrist Elis viduals expressing them could be counseled to refrain Essen-Möller and the plant breeder Herman Nilsson-Ehle from marrying. As an alternative, they could be steril- he created a network to promote eugenic consciousness ized. To protect the society at large, mentally disabled [12, 13]. The race biologists set out publishing popular persons or criminals could be sterilized or castrated booklets to educate the public and began lobbying for a against their will. Positive eugenics recommended various national institute for race biology. The painter Anders incentives to help the genetic elite to reproduce at least Zorn financed the Ethnic Type Fair (Rastypsutställningen), as fast as those less endowed. Saura Hereditas (2020) 157:48 Page 3 of 12

Sterilization of the feeble-minded, epileptics and the condition that the interest of the fund would be used asocial elements was a major goal of negative eugenics. to promote medical science. The Society decided that the Even though the evidence that the traits underlying interest of the fund would be used for “a comprehensive these characteristics were hereditary was often scant, the scientific examination of the mental and physical health of Nordic countries made in the 1930’s laws allowing the the Swedish-speaking population in Finland and of all cir- sterilization of the undesirables. The course of events cumstances, which might exercise influence on it. It was was essentially similar in each case, so that a short over- particularly pointed out that the significance of heredity view will suffice here. The German occupation of for the health of the people should be made a special ob- gave rise to an interlude in this development, ject of research” [22]. the most astonishing feature of which was that the laws Ossian Schauman and Jarl Hagelstam, professors of in- were repealed as late as in the 1970’s. I shall give a short ternal medicine and neurology, set with the zoologist overview of race biology and eugenics in Denmark, Harry Federley up a committee (Florinska kommissio- Finland, , Norway and Sweden. nen) to carry out the program outlined above. The com- mittee screened the occurrence of mental diseases, Denmark tuberculosis etc. In addition, anthropological measures, The anthropological committee (Den antropologiske housing conditions and nutritional status were recorded. Komité) was founded in 1905. The board members in- The anthropological survey was motivated through the cluded the police physician Søren Hansen, the pioneer success of the ethnic type fair in Sweden, but it was soon geneticist Wilhelm Johannsen, the statistician Harald abandoned. The pilot phase covered about 23.000 per- Westergaard and the psychiatrist August Wimmer [19]. sons. In addition, prizes were awarded to mothers with a The committee was a private society that obtained pub- healthy family background and at least four healthy and lic funding in 1920. It lobbied for eugenic legislation. strong children between the ages of 4 and 14 years [22]. Marriage restrictions were the first step in 1909–1913. Public Health in Swedish Finland (Samfundet Folkhäl- In 1912 Søren Hansen was elected member of the Per- san i Svenska Finland) was established in 1921 to pro- manent International Eugenics Committee in London. mote this program of positive eugenics. Schauman was Karl Kristian Steincke, a social democrat and minister of the first chairman, followed by Federley, by then the pro- justice in the 1920’s, formulated a race hygiene program fessor of genetics. The other members of the executive for the welfare state as a part of social reform. The con- board were professors of medicine and philanthropists. trol of reproduction was seen as a prerequisite to finance Folkhälsan created a network of community nurses to the welfare state. His proposal 1920 “The welfare state combat child mortality and campaigned against alcohol of the future” had eugenics as an integral part of social abuse, smoking and tuberculosis [21, 22]. reform. It was based on Darwinism, Mendelism and the In 1929 the government of Finland set up a committee degeneration hypothesis of the French psychiatrist Béné- to investigate whether compulsory sterilization of the dict Augustin Morel [11]. feeble minded etc. would be desirable. They had in mind The Danish Sterilization Commission of 1924 had the Danish preliminary law of 1929. The sterilization law three representatives of the Anthropological Committee, passed the parliament in 1935. It became, however, soon including Johannsen. He criticized the proceedings on apparent that the law was seldom enforced. Conse- theoretical, not practical grounds, pointing out that e.g. quently, a new and stricter law was put into effect in alcoholism is not hereditary. The Commission finished 1951. It was repealed in 1970 [23, 24]. Kajanoja [25] its job in 1926 and the preliminary sterilization law of sums up the anthropological studies made on Finns and 1929 was based on its work. It passed the parliament related peoples. with flying colors. Only the conservatives were skeptical. The preliminary law was replaced in 1934 and 1935 Iceland through a new one that made compulsory sterilization The professor of medicine Gudmundur Hannesson took possible for the mentally retarded until 1967 and for anthropologic measures on the Icelanders [26]. A criminals until 1972 [11, 20]. sterilization and abortion law was passed in 1937 in Iceland, the last one among the Nordic countries. People Finland diagnosed as feebleminded could be sterilized against The positive social aspect came first in Finland. The em- their consent [27]. The law was amended in 1975 [28]. phasis was preserving and furthering a language [21]. The majority of Finns spoke Finnish while Swedish was the Norway mother tongue of an influential minority. In 1911 Jenny The pharmacist Jon Alfred Mjøen prescribed eugenics to Florin donated 100000 Finnish marks to the Swedish Lit- cure disorders of the health of mind and body. He was erary Society in Finland (Svenska Litteratursällskapet)on politically affiliated with the liberal Venstre party. In Saura Hereditas (2020) 157:48 Page 4 of 12

1906 Mjøen not only founded and financed Winderen Norway in May 1945. The number of mental patients ster- Biological Laboratory (Winderen Biologiske Laboratorium) ilized following the introduction of the act rose at first in Kristiania () to study the effects of race crossing, i.e. sharply but evidently never reached the goal the between the Sámi and Nordic Norwegians and the inherit- had envisioned. The original law of the year 1934 was ance of intellectual capacity. He saw the , i.e. a abolished in 1977 [33]. takeover of East Asians, as an impending threat. Mjøen had contacts with , who founded in 1906 The Sweden German eugenics society (Deutsche Gesellschaft für Rassen- The Swedish Society for Race Biology had lobbied for hygienie). In 1908 Mjøen initiated the Norwegian eugenics the establishment of a Nobel Institute for Racial Biology program (Racebiologi og Fortplantninsgshygiene). In 1912 at the Karolinska Institute in Stockholm. These attempts The Permanent International Eugenics Committee in failed, but the Swedish Parliament founded the Swedish London used it as a model. Mjøen was appointed as a self- State Institute for Racial Biology (Statens Institut för evident member to the committee. He advocated negative Rasbiologi) in Uppsala in 1921 with Herman Lundborg measures such as the permanent segregation and as its director [13, 34]. The proposition to the Parlia- sterilizationoftheunfitaswellaspositiveonessuchasma- ment to found the Institute had been enthusiastically ternal insurance and propaganda for health and wholesome supported by virtually all politicians, including Hjalmar food along with prophylactic treatments to protect the un- Branting and Arvid Lindman, leaders of the Social born [29, 30]. Democratic and Conservative parties, respectively. The In 1914 Mjøen published a treatise titled Racie hy- Society published a jubilee volume [35] to commemorate giene, (Racehygiene,[31]). The university scientists Kris- the event. The book contains short entries on anthropol- tine Bonnevie and Otto Lous Mohr attacked his ideas as ogy, eugenics, cytology and genetics in Sweden and genetically unsound. Mohr had analyzed child mortality Finland. in relation to population increase (emigration from The Uppsala Institute opened its doors 1 January 1922 Norway was intense at that time). He criticized Mjøen’s with Lundborg as its head. It was then the only race view that western nations are dying because of an in- biology establishment in the world financed by a govern- crease of bad elements and advocated responsible birth ment [36] . It served as the model for The Kaiser control, which will lead to an automatic optimal breed- Wilhelm-Institut für Anthropologie, menschliche Erble- ing program. The animal breeder Christian Wriedt criti- hre und Eugenik founded in 1927 in Berlin. The Swedish cized Mjøen as well. Bonnevie and Mohr founded in Society for Race Biology and the State Institute remained 1919 Norsk forening for arvelighetsforskning (the Norwe- the only organizations of their kind in the Nordic coun- gian Society for Genetic Research) and kept Mjøen out. tries. In 1925 the Institute organized a Nordic meeting Undaunted, Mjøen initiated the Norwegian Consultative on race biology and anthropology that was attended by Committee for (Den norske konsultative some 30 participants [36]. komiteen for rasehygiene). The antagonism between Herman Lundborg was the director of the State Insti- Mjøen and the university geneticists was, in part, moti- tute from 1922 until 1935. He was mainly interested in vated through a competition for the meager financing race biology and published several popular booklets on available. Mjøen carried, however, the day: his ideas were the danger of degeneration. He argued that fecundity used as groundwork for eugenic legislation [30, 32]. was high in northern Sweden, i.e. a low-income area In 1919 Mjøen founded the journal The Nordic Race where race mixing was prevalent [37]. Olsson [38] had (Den Nordiske Race). It published papers mostly in Scan- in 1918 used a very extensive data set covering all of dinavian languages. J.A. Mjøen was the editor-in- Sweden to refute this argument. He showed that the cor- chief.and the publisher Winderen Laboratorium at Oslo; relation between fecundity with income was positive; the it was printed in Copenhagen. The editorial board in- middle class reproduced faster than the lower one. cluded an impressive list of luminaries such as Leonard Lundborg’s first magnum opus (quite literally, it shared Darwin, Charles Benedict Davenport, Herman Nilsson- the giant format of Retzius & Fürst 1902 [7] and Lund- Ehle, Henry Fairfield Osborn, Alfred Ploetz and Hans borg [10]) and was titled “The racial characters of the Virchow. Swedish nation” [34]; there was also an abridged version The Norwegian parliament passed in 1934 almost in German [39]. The book contains, in addition to exten- unanimously a law that allowed legal sterilization on sive statistics, wonderfully naïve photos. It is written in a eugenic and social grounds. Compulsory sterilization self-adulatory tone that is difficult to take seriously was allowed only in cases of severe mental deficiency. In today. It is no mean feat: the material is made up of 47, 1942 the Nazi puppet Quisling government pushed 387 men between 20 and 22 years old, some 50% of all through a sterilization act. It followed the severe German males in Sweden of that age cohort. Lundborg gave out model and remained in force until the liberation of a popular version with maps in Swedish [40], intended Saura Hereditas (2020) 157:48 Page 5 of 12

to be used in schools. In the above books Lundborg re- view in a nutshell. Dahlberg was one of the 23 signator- fers positively to the ideas of Nordic supremacy pro- ies of the eugenics manifesto of 1939 titled “Social Biol- posed by the linguist Rolf Nordenstreng (Nordenstreng ogy and Population Improvement” [51], a proclamation [41]. The Institute for Racial Biology in Uppsala took signed by leading geneticists 3 days before the onset of measures on some 100.000 people, mostly military con- World War II. This public declaration condemns scripts and minorities. but maintains a somewhat ambivalent attitude towards Herman Lundborg shared with Anders and Gustaf the social aspects of eugenics. Retzius the interest in measuring the Sámi. Lundborg & Wahlund [42] is a massive treatise on Sámi in the Race biology in Hereditas Retzius-Lundborg tradition. Like Lundborg & Linders As stated above, Hereditas was envisioned to cover ap- [34] it was well received. To mention an example, the plied race biology, in particular aspects deemed to be “of Dutch colonial authorities purchased six copies to Bata- utmost importance in the near future”. Robert Larsson, via (Jakarta). It was followed by Dahlberg & Wahlund the first editor-in-chief was an ardent supporter of eu- [43] that contained the photos and accessory material. genics [13, 17]. How did Hereditas carry into effect these The latter had the bulky format of the former, but it is expectations? The following review covers the contribu- much thinner. Here the authors repudiate physical tions to the subject, from race biology to human genetics ethnological race biology as obsolete in from the first issue of 1920 up to the end of race biology no uncertain words. The tenets of anthropological race in the early 1950’s. Let us take one author at a time and biology had begun to crumble on a wide front around in chronological order. This allows us to see how the au- 1934, e.g. [44, 45]. Lundborg had retired embittered in thors changed their views on a controversial subject that 1935 and Gunnar Dahlberg was his successor as the underwent a drastic paradigm change in the span of 30 head of the Institute for Race Biology. Lundborg’s swan years. song was Västerlandet i fara “The West in peril” [46], a demagogic booklet. Herman Lundborg The zoologist Nils von Hofsten, a board member of In 1920 Herman Lundborg was elected member of the the Institute of Racial Biology, reviewed in the capacity advisory board of Hereditas. He came out with a paper of a scientific expert on the State Medical Board (Social- on “The hereditary transmission of genotypical deaf- styrelsen) the applications for sterilization that were mutism” inthefirstissueofthefirstvolume,thesec- made legal through the law passed in 1934 until 1953. ond paper ever published in the journal [52]. Lundborg He reviewed and expanded the law of 1934 so that a had consulted the eminent human geneticist Wilhelm new law of 1941 was mainly his making. His credentials Weinberg in selecting a proper proband method [53]. as a geneticist are meager. He felt that it was the respon- Lundborg had another paper in the first volume as well sibility of the society to cut out the dead wood of the [54]. Using data on the incidence of tuberculosis in “rot in the stock” [13]. The sterilization law was repealed Sweden he developed an argument that the offspring of in 1976. race (i.e. Swede, Finn or Sámi) crosses are more suscep- A eugenicist streak had been present in the socialist tible to tuberculosis than the population at large. He used movement already in the early 1900’s. Arthur Engberg, a the plant breeding work of Oscar Hagem and Herman social democrat and newspaper editor had supported eu- Nilsson-Ehle as supporting evidence. In addition to north- genics and anti-Semitism in the 1910–20’s but in the ern Sweden, where the Swedes, Finns and Sámi inter- capacity of the minister of education and research he cut breed, he showed that levels of TBC in Stockholm and the financing of the State Institute in 1933. Engberg saw Gothenburg were as high as in the North. These big cities that Lundborg’s constant study trips to Lapland were a had attracted people from all parts of Sweden and from waste of money [47, 48]. The social democratic politician abroad, with race mixing as a result. The line of reasoning and (university chancellor) Östen Undén supported is roughly as follows: each race has developed a specific Gunnar Dahlberg as the successor of Lundborg instead genetic resistance to TBC. The genetic control of suscepti- of the eugenically oriented Torsten Sjögren [17]. bility is polygenic, organized in coadapted gene complexes Dahlberg had in 1922 started his career as Lundborg’s in race. In race crosses the resistance breaks down. The assistant at the Institute for Racial Biology. He oversaw rest of the paper is a routine description of eye and hair the change of the institute from one of ethnology to hu- color in the three races. Interestingly, he found out that man genetics. Dahlberg published e.g. on alcoholism and red hair is associated with diseases and criminal behavior. diseases as a social problem and debunked doctrines of He drew conclusions on the basis of raw data without the race in popular books such as Dahlberg [49]. In particu- benefit of a statistical analysis. lar “Race reason and rubbish” [50] was widely read and Lundborg [55] developed the above hypothesis on the translated into several languages. It contains the modern deleterious effects of interbreeding between races further. Saura Hereditas (2020) 157:48 Page 6 of 12

He started from the observation that people with a mixed bridge history and genetics at a time when the learned dis- parentage are taller but argued that they are disease- course underwent a dramatic change? prone. Mixing races results in a chaotic genetic constitu- In 1921 Nilsson [62] explains the fall of Rome as a re- tion “Genchaos” with an unbalanced phenotype as a con- sult of unfettered bastardizing between races. He was sequence. Here he has extended the study to cover the certainly acquainted with an immensely influential book entire European continent. His examples cover , of the time. Oswald Spengler argued in Der Untergang Germans and Russians. des Abendlandes (The decline of the West) that civiliza- He [56] measured eye and hair color, and cepthalic tions are superorganisms each with a predictable and and facial indices in the population of the northernmost limited lifespan. The West (i.e. the Greco-Roman part of Sweden. The Swedes and Finns had light eyes civilization) had fallen once and the second fall is com- and fair hair, while the Sámi had dark hues. The Swedes ing near. Spengler had used the word “race” copiously, and the Sámi had extremes in head dimensions with the but not in a biological sense. As told above, Lundborg Finns intermediate between the two. There had been and Mjøen had promoted the deleterious biological ef- gene flow from the Sámi to the Finns; when the former fect of race mixing. abandon their nomadic way of life, they adopt Finnish Bengtsson [63] and I [37] have described the contents customs. of Nilsson [62] so that a short rehearsal only will suffice Lundborg’s ideas represent mainstream race biology of here. Inbreeding within a race was the rule in olden the 1920’s. E.g. Jon Alfred Mjøen and others voiced simi- times. Later on, Roman upper classes did not propagate lar concerns [57]. – In addition to the above papers that their kind fast enough with the result that the scum of we now may consider to represent rather unrestrained the earth populated the capital. The empire gave rise to , Lundborg had a short report [58]on unlimited bastardizing; mixing races lowers the fitness of the sex-linked inheritance of ichthyosis simplex. The the better one. Hybrids may have been intelligent but condition is characterized with rough and scaly skin; of they lacked moral stamina. The result was chaos. Evi- the multiple forms of the disease this is the mildest one. dently Nilsson had read Lundborg [54, 55] who used an Lundborg used church records to demonstrate how this identical terminology in a medical context. Herman form of ichthyosis is inherited. Nilsson-Ehle a friend of Nilsson [64], evidently told him what was newest new in Hereditas. Nilsson expanded Emanuel Bergman the above ideas in a booklet published by the Society for In the first volume of Hereditas the Swedish physician Racial Biology [65]. ’ Bergman described a family with hereditary tremor [59]. Nilsson s second contribution to Hereditas [63]isan This is pure human genetics. about-face [37]. He has clearly intended to write an essay rather than a scientific treatise. He points out that dog breeders mix races to obtain desirable characteristics. Halfdan Bryn Nilsson notes briefly that he [62] had argued that the Bryn was a military physician and anthropologist in chaos of non-complementary genes brought down the Trøndelag in central Norway. In the first volume of Roman Empire. Hereditas Bryn [60] published the results of extensive Here Nilsson demonstrates that the ancient civiliza- observations on eye color and cephalic index in Selbu tions were created and borne by mixed peoples. His ex- and Tydal, two nearby but isolated municipalities. He amples include the Egyptians, Babylonians, Assyrians, tried to find out the inheritance of these above traits and Greeks, citizens of the Roman Republic etc. The Jews, in did not discuss the results in an anthropological context. particular, are a gifted people with multiple origins. An Later on, he may have been somewhat carried away with initial chaos will give rise to a number of biotypes: some a nationalistic sense of superiority (he argued that Cro- are definitely positive and persist while some others will Magnon people still lived in parts of Norway etc.) [61]. disappear in the course of genetic segregation and selec- tion. Indo-Europeans were a destructive element that Martin P:son Nilsson was domesticated through interbreeding with the older Nilsson was the professor of ancient Greek, classical Europeans. All peoples of Europe and the Near East des- archeology and ancient history at Lund. His two contri- cend from this process. butions to Hereditas the first in 1921 and the second Nilsson was an outsider in a genetics journal. This one in 1939 [62, 63] represent a remarkable interdisciplin- gave him an opportunity to see how times and values ary approach intended to give a genetic explanation to his- had changed in the span of less than twenty years. In tory. Natural sciences and humanities are often seen to 1921 he [62] wrote at a time when physical race biology represent two separate cultures, characterized by mutual was cutting-edge science, backed through a political misunderstanding and controversies. How did Nilsson consensus. Sterilization laws would follow in due time. Saura Hereditas (2020) 157:48 Page 7 of 12

As we shall see, eugenics changed direction in the 1930’s The reason why I take up Turesson in the company of from sterilization towards a social policy of the welfare race biologists is that he furthered typological thinking state. Anthropological race biology was increasingly seen to biology and Hereditas at a time when population gen- as an aberration. Hitler’s rise to power had led to vulgar etics was on the rise among human geneticists. Nilsson’s and unrestrained doctrines of race that in the Nordic [61] and Lundborg’s[55, 56] reasoning comes close to countries were seen to represent a travesty of science. Turesson’s. Turesson had, however, based his conclusions Nilsson’s essay of 1939 [63] was printed in the June of on experimental evidence and on plants. Unlike animals, the last summer of peace in the long shadow of the plants control the process of fertilization through breeding impending war that would engulf four of the five Nordic systems, which range from complete apomixis (each plant countries. makes identical copies or clones of itself) to complete out- breeding imposed by dioicy or self-incompatibility sys- Søren Hansen tems. Population biologists have often difficulties in Hansen was a leading Danish anthropologist and a found- understanding that plants need not behave as members of ing member of the Permanent International Eugenics an ideal population. Committee. He had a single contribution to Hereditas in 1922 [66], a study on the inheritance of dementia praecox Kristine Bonnevie (schizophrenia). He uses European royal houses to illus- In 1912 Bonnevie was appointed professor of zoology at the trate the contribution of heredity vs. environment and University of Kristiania (Oslo). She had studied widely ponders the plausibility of different alternatives. It repre- abroad, namely in the laboratories of Arnold Lang in sents human genetics pure and simple without eugenic Zürich, in Würzburg and Edmund Wilson overtones. in New York. It is difficult to imagine more outstanding hosts. Lang hed developed cytological techniques, Boveri and Wilson’s student Sutton showed that genes reside in Kaarlo Hildén chromosomes. Bonnevie specialized in the cytology of mar- Hildén was the professor of geography at the Helsinki ine invertebrates. In 1916 she founded the Institute for School of Economics. His credentials as an anthropolo- Genetic Research to study human genetics at the University gist included an expedition to the Altai mountains in of Kristiania [72]. Bonnevie collaborated with the Drosoph- Siberia. In 1921 he undertook to study the isolated and ila geneticist Otto Lous Mohr, the professor of anatomy. inbred Swedish speaking population on the island of Bonnevie had three papers in Hereditas [73–75]. She Ruhnu (Runö), about equidistant between Estonia and quantified the differences underlying fingerprint patterns Latvia in the Gulf of Riga. Using observations and and attempted to determine how they are inherited. In church records he analyzed the inheritance of the shape the second paper she tried to find out whether the fin- of the earlobe Hildén [67], head dimensions [68] and the gerprint pattern was associated with IQ among school- shape of the nose [69]. These three papers in Hereditas children. The patterns were used in the diagnosis of describe the inheritance of characters relevant to metal deficiency. She noted that fingerprints had been anthropometrics. used in ethnographic contexts but did not take up race biology. The third is about the relation of left- Göte Turesson handedness to fingerprints. Göte Turesson is certainly one of the foremost authors in Hereditas. The reason that he here has fallen among Gert Bonnier human race biologists is certainly not his making. Turesson A Drosophila geneticist, Bonnier studied theoretical as- made truly pioneering contributions to ecological genetics, pects of human genetics; something that may have published in Hereditas. Herman Nilsson-Ehle supervised helped him to establish the animal breeding institute at his PhD study at Åkarp and Lund. Turesson’s basic ap- Wiad in the 1930’s. He was the first professor of genetics proach is seen in ’s influential textbook [70]. of the University of Stockholm. Baur was a friend of Nilsson-Ehle and Scandinavian nature. Bonnier [76] criticized in Acta Zoologica (the journal Turesson observed variation in common plants with of Stockholm zoologists) that the association of hetero- different breeding systems both in the wild and under zygosity and TBC proposed by Lundborg in 1920 and cultivation, including transplantation experiments. Some 1921 [55, 56] in Hereditas. In Hereditas he used French, plants remained constant while some responded to the the language of mathematics, in defining the application new environment. He partitioned the variation into a of statistics in biology [77]; a theme that he developed phenotypic and genotypic component and he coined in later on in a textbook. Even though human cytogenetics 1922 the term ecotype to describe the genotypic re- was at an elementary stage in the 1930’s, he described a sponse to the habitat [71]. case of heritable sex-mosaicism in Hereditas [78]. Saura Hereditas (2020) 157:48 Page 8 of 12

Professor Solomon Levit of Moscow criticized his statis- Lännart Ribbing tics and consequently Bonnier [79] published an apology Ribbing, a specialist in the comparative anatomy of ver- and correction. This did not reach Dr. Levit; Lysenko tebrates, made an entry to race biology: he took mea- had smeared him as traitor and he was shot in early sures on Icelanders. He was a member of first executive 1938. board of the Mendelian Society and presided at the first meetings [16]. In 1922 he was appointed professor of anatomy at the Royal Swedish Academy of Fine Arts. In Gunnar Dahlberg addition, Ribbing won acclaim as a poet. He was one of Dahlberg had studied medicine at Uppsala. As men- the few who published in French in Hereditas. tioned earlier, he was appointed Lundborg’s assistant at Ribbing had a single contribution to Hereditas [88]. It the State Institute for racial biology in 1922 and, in represents skull measurement or craniology, based on 1935, was elected Lundborg’s successor as the head of three skulls dug out of the earth at Vellinge in south- the institute. He understood that both statistics and western Scania. The oldest of the skulls hails from Stone population genetics showed the hollowness of race biol- Age, the next from Iron Age while the third is modern. ogy. Consequently, he changed the name of the Institute This modest sample of three is meticulously photo- to one of human genetics and racial biology. graphed and all pertinent parameters are given. In his first paper in Hereditas in 1923 Dahlberg [80] Why was a contribution like this included in a journal used statistics to differentiate between mono- and dizyg- otherwise dedicated to the branch of science founded by otic twins in two extensive data sets, one collected from Friar ? Ribbing argues that the skulls are Germany by Weinberg and the other by Bonnevie in remarkably similar. Consequently, a morphologically Norway. uniform human population has inhabited Vellinge since In 1930 Dahlberg had two papers in Hereditas [81, 82]. the days of the first one: “an intelligent person repre- The first introduced a new statistical method in treating senting a distinguished culture” [88]. family data. It drew forth a response from Wilhelm Weinberg [53, 83] in Hereditas. His second paper [82]isan Sven Wahlund analysis on the effect of inbreeding in different modes of in- Wahlund, a statistician at the State Institute for Racial heritance. Dahlberg showed here, as in his later papers in Biology at Uppsala, published with Lundborg and Dahlberg Hereditas [84] a good grasp of both statistics and popula- on Sámi [42, 43]) but he had papers on plant breeding in tion genetics. Hereditas as well. He is the author of one of the truly sem- inal papers ever published in Hereditas: Wahlund 1928 [89], in which he describes the Wahlund effect,aningredi- Victor Berglund ent in standard texts on population genetics. This arguably Berglund was a Swedish physician, painter and sculptor most famous Swedish contribution to population genetics had a contribution on the occurrence of hypotrichosis states that as long as there is allele frequency variation (weak hair growth) within a family that was published among subdivisions of a population, the genotype frequen- posthumously in 1924 [85]. cies of the population will exhibit a deficiency of heterozy- gotes and an excess of homozygotes relative to Hardy- Weinberg expectations. Jon Alfred Mjøen In 1929 Wahlund [90] demonstrated a potential use As stated above, the eugenicist and race biologist Mjøen for the enormous amount of data stored at the Institute may not have earned kudos as a geneticist, but he suc- for Racial Biology. Lundborg and Linders [34] had ar- ceeded in having papers published not only in Hereditas gued that anthropological measures are associated with but also in e.g. Zeitschrift für induktive Abstammungs- social strata; the ones better off are taller and have a und Vererbungslehre, in spite of,or rather because of, higher cephalic index than the population at large, while the rather unusual subject, namely the inheritance of German authors seemed to have found evidence that the musical talent [86]. He used multiple approaches in the “Germanic” characteristics are selected against in cities. study: he sent questionnaires wide and far, observed dir- Using a sample of 677 students in among the 47.887 ectly the level of perception and assessed the contribu- conscripts arranged in order by Lundborg & Linders tion of the parents on musical talent in large families. [34], Wahlund compared the means of different body The results were straightforward. Untalented parents parts among students and the general population of have seldom talented children, while talented parents Sweden. The students were, on an average, 3 cm taller never have untalented offspring. Fritjof Mjöen, J.A. and differed significantly also from their compatriots of Mjøen’s son, extended these observations to include the the same age in the width of the zygomatic arch etc. perception of pitch [87]. Interestingly, the results were neither correlated to the Saura Hereditas (2020) 157:48 Page 9 of 12

geographical origin of the students nor to the social sta- time at the Grini concentration camp). The new depart- tus of their parents. ment folded when the university was shut down in late Wahlund notes that the results are apparently associ- 1943 [98]. ated with the upward social mobility of the students. What is cause and what is effect is left open. Wahlund Jan Arvid Böök warns against generalizing the results to other popula- BöökmovedfromLundtoUppsala,wherehewas tions; there is simply no reference material anywhere. appointed head of the State Institute of Medical Genetics and Racial Biology in 1957 following Dahlberg’s retirement. Oluf Thomsen As told above, Dahlberg had directed the State Institute to Thomsen was the professor of pathology at the Univer- one of human genetics away from the race biology of sity of Copenhagen. He was the driving force in the es- Lundborg. Böök cut the last ties to race biology: in 1959 tablishment of the Department of Human genetics at the State Institute was incorporated into the University of Copenhagen. His first contribution to Hereditas [91] was Uppsala as the Department of Human Genetics [99]. a description of a hereditary growth anomaly of the Böök had two papers in Hereditas in 1948 [100, 101]. thumb. He shifted his attention to the inheritance of Both deal with the effect of cousin marriages, so that the blood groups [92, 93] and several others. latter takes up the inheritance of clubfoot, including the Thomsen wrote popular presentations of genetics. mutation rate and twin studies. They had titles such as Human heredity (Menneskets Arvelighedsforhold). The first editions contained much The Eighth International Congress of Genetics eugenics. His outlook changed as years passed and he The Stockholm International Congress of Genetics was was actively studying blood groups and he became held in 1948. The proceedings produced a flurry of pa- acquainted with population genetics. An edition of 1934 pers, published as supplement 35S in Hereditas a year [94] has chapters on eugenics and human races. As for later. There is a single paper on anthropological race eugenics he points out that there is no need to do any- biology: Corrado Gini of Rome presented a treatise on thing about dominant traits, since they will take care of the physical assimilation of Albanians to a host popula- themselves. Anything with recessive traits will be inef- tion in Calabria, Italy [102]. The Calabrians were short fective. Thomsen advocates “passivity” in the enforce- and dolichocephalic, while the Albanians were tall and ment of sterilization laws. He downplays races; points brachycephalic. Gini gives comparisons among several out that they are not uniform with regard to any alleles other physical characteristics as well. etc. He shows that the Jews are not a race and that the There were four non-Nordic papers on human genet- races of Europe are, if anything, mixed. ics, namely JBS Haldane on the mutation rate of human genes [103], Lionel Sharples Penrose on palmar patterns Torsten Sjögren in relation to mongolism (Down’s syndrome) [104], Lau- Sjögren was employed at the Institute for Racial Biology rence H. Snyder on linkage relationships of sickle-cell in 1925. He had an extensive treatise on Tay-Sachs dis- anemia [105] and Alexander S. Wiener on Rh blood ease ((juvenile amaurotic idiocy) in 1931 in Hereditas types [106]. [95] as well as a contribution to the Eighth International Congress of Genetics in 1949 [96]. He was a serious con- Tage Kemp tender for the chair of director vacated by Lundborg. A Tage Kemp studied medicine at Copenhagen with Oluf psychiatrist, he believed in the efficacy of sterilization Thomsen as supervisor. He visited ’s already in childhood [17]. He has gained some notoriety Eugenics Record Office at Cold Spring Harbor. Davenport as a member of eugenic organizations that had a racist was the leader of the American eugenics movement. In tinge [97]. Sjögren was the professor of psychiatry at 1932 Kemp participated in the Third International Eugenics Karolinska Institutet in Stockholm from 1945 to 1961. Congress in New York with a paper on prostitution and heredity. He studied chromosomes and mouse genetics and Thordar Quelprud had a paper on the latter subject in Hereditas. The Univer- Quelprud, a student of Bonnevie, described a Drosophila sity Institute of Human Genetics and Eugenics was estab- mutant in Hereditas. He studied anatomy in Germany, lished in 1938 on the initiative of Oluf Thomsen. He had where he became a naive national socialist Even though contacted the Rockefeller Foundation in 1934 and obtained he was not a PhD, the Quisling administration made 250.000 Danish Crowns + 110.000 Crowns as a grant, the him docent in 1942 and he created a Department of interest of which would be used to maintain the institute. Hereditary Biology at the , located be- The Danish state would be responsible for the personnel tween the premises of Bonnevie and the anatomist and and day-to-day costs. Kemp was the head of the institute anthropologist Emil Kristian Schreiner (who had done from 1938 on; ten years later he was installed as professor Saura Hereditas (2020) 157:48 Page 10 of 12

of genetics and eugenics at the Medical Faculty of the Uni- ten editions and was translated into several languages. versity. Kemp abhorred the propagandizing and pseudosci- The book is strong on cytogenetics as well as plant entific Nazi race hygiene. He even started criticizing and animal breeding. It has photos of Europeans be- eugenics, a stance recommended by the Rockefeller Foun- longing to different races presented in the traditional dation in the late 1930’s[11]. typological fashion, including the Sámi as a kind of Kemp had a contribution to the Eighth International outgroup. Müntzing condemned racism in no uncer- Congress of Genetics in Stockholm titled “The rise of tain terms. He changed eugenics of the first editions human genetics” [107]. It is a short history of the subject to genetic counseling later on. Mogens Westergaard with a particular emphasis on the development in of Copenhagen published also in 1953 a general text- Copenhagen. book of genetics in Danish: Arvelighedslæren.He shared the general outlook with Müntzing and in- Nils von Hofsten cluded some race biology [112]. As told above, von Hofsten had the task of reviewing the applications for sterilization that were sent to the State Conclusions Medical Board of Sweden. He was mainly responsible Biological anthropology was effectively dead as a branch for the new law of 1941. He evidently never did original of genetics in the late 1930’s. To mention an example, in research on genetics but wrote popular papers on the US and his students showed that race sterilization in Swedish in the 1930’s and 1940’s. He had categories were overlapping, ill-defined and historically written a textbook of genetics in 1919 that (in spite of inconsistent. They showed that the head form of immi- some howling errors) is a balanced treatment [108]. As grants changed from one generation to the next with an outsider, he had no ax to grind. He presented eugenic changed conditions of life; in other words it was subject literature and warned that the books of Mjøen and, in to the influence of the environment [112]. The horrors particular, the British eugenicist Caleb W. Saleeby, of the Second World War finished off what little may should be read critically. In addition, von Hofsten criti- have remained. The First International Congress of Hu- cized the extreme views of American eugenicists. man Genetics held in Copenhagen in 1956 did not men- von Hofsten had one paper on the subject of eugenics tion things such as race, or nation, the topic of interest in 1951 in Hereditas [109]: “The genetic effect of nega- was the human race or mankind. tive selection in man”. The paper extends over more Laws allowing sterilization on eugenic grounds were than 100 pages of curves, graphs and illustrations. In ef- enacted in Denmark, Finland, Iceland, Norway and fect it is a rehearsal. To give an example, HTJ Norton Sweden in a course of a few years in the 1930’s. The had worked much of it out in an appendix to a book on sterilization policies are now seen as a major human tra- mimicry in butterflies in 1915 [110]. In retrospect, one gedy: compensation is paid to its victims. Interestigly, wonders why this overlong paper was ever written. Von there is only a single paper discussing the justification of Hofsten had seen that “the sterilization policy had been negative eugenics in Hereditas [109]. It represents a be- the subject of a lively interest at the International Con- lated defense of a policy that had lost its adherents gress of Genetics in 1948”. Consequently, he wished to within the community of Nordic geneticists. demonstrate that sterilization had a firm scientific foun- In the 1950’s the university departments changed dation: genetic ills can express themselves in many ways names. Human genetics had an advisory function that and one must know how to get rid of them. guided the work of hospitals and doctors. It took some 15 years before the sterilization laws were abolished. Arne Müntzing This put an end to this advisory function as well. It had Müntzing has a secure place in Hereditas’ Hall of Fame. already earlier been supplanted by prenatal diagnosis In the 1930’s he published a series of papers in our and a research program differentiated to cover a multi- journal that were certainly ground-breaking, e.g. tude of genetic diseases. [111]. He had crossed two diploid hemp-nettles, As mentioned in the beginning, race is now hardly a Galeopsis speciosa with G. pubescens and obtained concern for scientists [1]). It remains, however, a virtu- tetraploid offspring that clearly represented a well- ally explosive political issue. August names such as known tetraploid taxon, namely G. tetrahit.Inshort, Linnaeus and Ronald A. Fisher are removed from public Müntzing had shown how a plant species had origi- consciousness [113] for their sins. The former had at- nated. He became the respected doyen of Nordic gen- tributed negative characteristics to nonwhite populations etics for years to come, not least as the editor-in- and the latter had championed eugenics. chief of Hereditas through 1955–1977. Hereditas was envisioned to treat race biology as a Müntzing wrote an influential textbook of genetics, major branch of applied genetics, on par with plant Genetic Research (Ärftlighetsforsking). It went through breeding in importance. The contents of Hereditas show Saura Hereditas (2020) 157:48 Page 11 of 12

how the ideas of race gave way to human genetics. 22. Schauman O. In: Lundborg HA, Runnström J, editors. The Swedish nation in Hereditas has a long record of publishing milestones in word and picture, together with short summaries of the contributions mades by Swedes within the fields of anthropology, race-biology, genetics human genetics such as the demonstration of the human and eugenics; a jubilee book given out, with the cooperation of experts chromosome number by Tjio and Levan [114] and the commissioned by the Swedish society for race-biology 1921. Eugenic work identification of all human chromosomes [115]. in Swedish Finland. Uppsala: Hasse W. Tullberg Co.; 1921. p. 89–94. 23. Hietala M. From race hygiene to sterilization. In: Broberg G, Roll-Hansen N, Author’s contributions editors. The eugenics movement in Finland, Eugenics and the welfare state. The author reads and approved the final manuscript. The author is the sole Sterilization policy in Denmark, Sweden, Norway and Finland: East Lansing, – contributor. Michigan State University Press; 1996. p. 195 258. 24. Hemminki E, Rasimus A, Forsius E. Sterilization in Finland: from eugenics to contraception. Soc Sci Med. 1997;45:1875–84. Funding 25. Kajanoja P. A study in the morphology of the Finns and its relation to the Open Access funding provided by University of Umea. settlement of Finland. Ann Acad Sci Fennicae Ser A V. 1971;146:1–86. 26. Körpermasze HG, der Isländer K. Ein Beitrag zur Anthropologie Islands. Competing interests Reykjavik: Árbók Háskóla Islands (Suppl); 1925. The author declares that he has no competing interests. The author is the 27. Hauge M. What has happened to eugenics? Eugen Rev. 1965;56:203–5. sole contributor. 28. Eydal GB, Olafsson S. Social and family policy. The case of Iceland. In: Third report for the project Welfare policy and employment in the context of Received: 3 November 2020 Accepted: 13 November 2020 family change. CiteSeer˟ www.york.ac.uk; 2003. 29. Hodson CBS. Eugenics in Norway. Eugen Rev. 1935;27:41–4. 30. Kyllingstad JR. Measuring the . Cambridge: Open book References publishers; 2014. 1. Nyborg H. Race as a social construct. Psych. 2019;1:139–65. 31. Mjøen JA. Racehygiene. Kristiania: Jacob Dybwads Forlag; 1915. (In 2. Basu A, Sarker-Roy N, Majumder PP. Genomic reconctruction of the history Norwegian). of extant population of India reveals five distinct ancestral components and 32. Hviid Nielsen T, Monsen A, Tennøe T. Livets tre og kodenes kode. Oslo: a complex structure. Proc Natl Acad Sci U S A. 2016;113:1594–9. Gyldendal; 2000. (In Norwegian). 3. Retzius A. Anders Retzii samlade skrifter af ethnologiskt innehåll. Stockholm: 33. Haave P. Sterilization under the . The case of Norway. Int J Ment Svenska Läkaresällskapet; 1864. (In Swedish). Health. 2007;36:45–57. 4. von Düben G. Om Lappland och lapparne, företrädesvis de svenske. Stockholm: Norstedt; 1873. (In Swedish). 34. Lundborg H, Linders FJ. The racial characters of the Swedish nation. 5. Kyllingstad JR. Measurng the Master Race. In: Physical anthropology in Uppsala: Almqvist & Wiksell; 1926. Norway 189–1945. Cambridge: Open Book Publishers; 2004. 35. Lundborg HA, Runnström, J, editors. The Swedish nation in word and 6. Retzius G. Finska kranier jämte några andra natur- och litteratur-studier picture, together with short summaries of the contributions mades by inom andra områden af Finsk Antropologi. Stockholm: Samson & Wallin; Swedes within the fields of anthropology, race-biology, genetics and 1878. (In Swedish). eugenics; a jubilee book given out, with the cooperation of experts 7. Retzius G, Fürst CM. Anthropologia suecica. Beiträge zur Anthropologie der commissioned by the Swedish society for race-biology; Uppsala, Hasse W. Schweden. Stockholm: Aftonbladets tryckeri; 1902. Tullberg Co. 1922. 8. Montelius O. In: Lundborg HA, Runnström J, editors. The Swedish nation in 36. Broberg G. 2002. Statlig rasforskning En historik över rasbiologiska institutet, word and picture, together with short summaries of the contributions vol. 4. Lund: Ugglan; 2002. (In Swedish). – mades by Swedes within the fields of anthropology, race-biology, genetics 37. Saura A. 2014 A tale of two papers. Hereditas. 2014;151:119 22. and eugenics; a jubilee book given out, with the cooperation of experts 38. Olsson JON. Statistiska studier över den äktenskapliga fruktsamheten med commissioned by the Swedish society for race-biology The immigration of särskild hänsyn till dess beroende av den sociala ställningen. Uppsala: NJ our forefathers to the north. Uppsala: Hasse W. Tullberg Co; 1921. p. 7–11. Gompert; 1918. (In Swedish). 9. Lundborg H. Die progressive Myoklonus-Epileåsie (Unerricht’s Myoklonie). 39. Lundborg H, Krauss W, Linders FJ. Rassenkunde des schwedischen Volkes. Uppsala: Almqvist & Wiksell; 1903. Jena: G Springer; 1928. 10. Lundborg H. Medizinisch-biologische Familienforschungen innerhalb eines 40. Lundborg H. Svensk raskunskap. In: Folkupplaga med text, kartor, diagram 2232köpfigen Bauerngeschlechtes in Schweden (Provinz Blekinge). Jena: och planscher. Uppsala: Almqvist & Wiksell; 1927. (In Swedish). Gustav Fischer; 1913. 41. Nordenstreng R. Europas människoraser och folkslag. Stockholm: Norstedts; 11. Koch L. Racehygiejne i Danmark 1920-56. Copenhagen: Gyldendal; 2000. (In 1917. (In Swedish). Danish). 42. Lundborg H, Wahlund S. The race biology of the Swedish Lapps. Uppsala: 12. Gustafsson Å. Herman Nilsson-Ehle, minnesteckning 175; levnadsteckningar Almqvist & Wiksell; 1932. over K. Svenska Vetenskapsakedemiens ledamöter. Uppsala: Almqvist & 43. Dahlberg G, Wahlund S. The race biology of the Swedish Lapps Pt. 2. Wiksell; 1972. p. 279–93. (In Swedish). Anthropogenic Survey. Uppsala: Almqvist & Wiksell; 1941. 13. Björkman M, stammen D a, von Hofsten N. eugeniken och 44. Backman G. Människoraserna och moderna rasproblem. Stockholm: steriliseringslagarna 1909-1963. Lund: Arkiv; 2011. (In Swedish). Bonniers; 1934. (In Swedish). 14. Petersson H. Nationalstaten och arkeologin. 100 år av neolitisk 45. Thomsen O. “Menneskets Arvelighedsforhold”, (Kultur og Videnskab). Poul forskningshistoria och dess relationer till samhällspolitiska förändringar. Branner: Copenhagen; 1934. (in Danish). Gothenburg: Gotarc B, No. 36, Dept. of archaeology; 2005. (In Swedish). 46. Lundborg H. Västerlandet i fara. Befolkningsfrågor i biologisk och 15. Bengtsson B-O. Genetik och politik. Stockholm: Norstedts; 1999. (In rasbiologisk belysning. Gothenburg: Ernst v. Hansson; 1934. (In Swedish). Swedish). 47. Blomqvist H. Nation, ras och civilisation/Svensk arbetarrörelse före nazismen. 16. Höglund M, Bengtsson BO. The origin of the Mendelian Society in Lund Stockholm: Carlsson; 2006. (In Swedish). and the start of Hereditas. Hereditas. 2014;151:110–4. 48. Hagerman M, Herman K. Rasbiologen Herman Lundborgs gåta. Stockholm: 17. Tunlid A. Ärflighetsforskingens gränser. Lund: Ugglan; 2004. (In Swedish). Norstedts; 2015. (In Swedish). 18. Porter TM. 2018. Genetics in the madhouse: the unknown history of human 49. Dahlberg G. Arv och ras. Solna: Kooperativa förbundet; 1940. (In Swedish). heredity. Princeton: Princeton University Press; 2018. 50. Dahlberg G. Race, reason and rubbish. London: G. Allen & Unwin Ltd.; 1942. 19. Wimmer A. 1929. Sindssygesdommenes Arvegang og raceforbedrende 51. Crew FAE, Darlington CD. Haldane JBS and 20 others. Social biology and Bestræbelser. Copenhagen: Levin & Munksgaard; 1929. (In Danish). population improvement. Nature. 1939;144:521–2. 20. Kemp T. Genetic-hygienic experiences in Denmark in recent years. Eugen 52. Lundborg H. The hereditary transmission of genotypical deaf-mutism. Rev. 1957;49:11–8. Hereditas. 1920a;1:35–40. 21. Saura A. Sata vuotta perinnöllisyystiedettä Suomessa. Helsinki: Vanamo; 53. Sperlich D, Früh D, Weinberg W. Der zweite Vater des Hardy-Weinberg- 2018. (in Finnish). Gestzes. Freiburg im Breisgau: Basilisken-Presse; 2014. Saura Hereditas (2020) 157:48 Page 12 of 12

54. Lundborg H. Rassen- und Gesellschaftsprobleme in genetischer und 90. Wahlund S. Sozialer Aufstieg und anthropologische Auslese. Hereditas. 1929; medizinischer Beleuchtung. Hereditas. 1920b;1:135–77. 12:71–108. 55. Lundborg H. Rassenmischung – vermehrte Heterozygotie (Genchaos) – 91. Thomsen O. Hereditary growth anomaly of the thumb. Hereditas. 1928;10: Konstitutionsveränderungen – Habitus asthenicus sive paralyticus (Zunahme 261–73. der Körpergrösse usw.) – Tuberkulose. Eine Ursachenkette. Hereditas. 1921;2: 92. Thomsen O. Die Erblichkeitsverhältnisse der menschlichen Blutgruppen: Mit 77–8. besonderem Hinblick auf zwei “neue” A’ und A’B genannte Blutgruppen. 56. Lundborg H. Racial structure of the Finns of the Northernmost part of Hereditas. 1930;13:121–3. Sweden. A short analysis and a preliminary survey. Hereditas. 1923;4:125–41. 93. Wellisch S, Thomsen O. Über die Vier-Gen-Hypothese Thomsens. Hereditas. 57. Provine WB. Scientists and the biology of race crossing. Science. 1973;182: 1930;13:121–63. 790–6. 94. Thomsen O. Menneskets arvelighedsforhold Copenhagen. Poul Branner: 58. Lundborg H. Geschlechtsgebundene Vererbung von ichthyosis simplex Copenhagen; 1935. (In Danish). (vulgaris) in einer schwedischen Bauernsippe. Hereditas. 1927;9:45–8. 95. Sjögren T. Die juvenile amaurotische Idiotie. Klinische und 59. Bergman E. A family with hereditary (genotypical) tremor. Hereditas. 1920;1: erblichkeitsmedizinische Untersuchungen. Hereditas. 1931;14:197–425. 98–106. 96. Sjögren T. A genetic-psychiatric study of a Swedish rural population. 60. Bryn H. Researches into anthropological heredity. Hereditas. 1920;1:186–212. Hereditas. 1949;35S:429–45. 61. Bryn H. Der nordische Mensch: die Merkmale der nordischen Rasse mit 97. Kühl S. Die Internationale der Rassisten. In: Der Aufstieg und Niedergang besonderer Berücksichtigung der rassiscen Verhältnisse Norwegens. der internationalen Bewegung für Eugenik und Rassenhygiene im München: J F Lehmann; 1929. zwanzigsten Jahrhundert. Frankfurt am Main: Campus; 1997. 62. Nilsson MP. The race problem of the Roman Empire. Hereditas. 1921;2:370–90. 98. Vogt Y. Udugelige naziprofessorer. Oslo: Apollon; 2007. (In Norwegian). 63. Nilsson MP. Über Genetik und Geschichte. Hereditas. 1939;25:210–23. 99. Lindroth S. A history of 1477-1977. Uppsala: Almqvist & 64. Bengtsson BO. Strange history: the fall of Rome explaned in Hereditas. Wiksell; 1976. Hereditas. 2014;151:132–59. 100. Böök JA. The frequency of cousin marriages in three North Swedish 65. Nilsson MP. Rasblandningarnas omfång och betydelse i det romerska parishes. Hereditas. 1948;34:252–5. kejsarriket. Sven Sällsk Rashygien Skriftser. 1923;11:1–25 (In Swedish). 101. Böök JA. A contribution to the genetics of congenital clubfoot. Hereditas. 66. Hansen S. Heredity in the case of dementia praecox and related diseases. 1948b;34:289–300. Hereditas. 1922;3:114–23. 102. Gini C. The physical assimilation of the descendants of immigrants. – 67. Hildén K. Über die Form des Ohrläppchens beim Menschen und ihre Hereditas. 1949;35S:234 43. Abhängigkeit von Erbanlagen. Hereditas. 1922;3:351–7. 103. Haldane JBS. The rate of mutation of human genes. Hereditas. 1949;35S: – 68. Hildén K. Zur Kenntnis der menschlichen Kopfform in genetischer Hinsicht. 267 73. Hereditas. 1925;6:127–46. 104. Penrose LS. Familial studies on palmar patterns in relation to mongolism. – 69. Hildén K. Zur Kenntnis der Erbfaktoren des menshlichen Nasenform. Hereditas. 1949;35S:412 6. Hereditas. 1920;13:87–106. 105. Snyder LH. Studies on human inheritance XXXV. The linkage relationships of – 70. Baur E. Einführung in die experimentalle Vererbungslehre. Berlin: sickle cell anemia. Hereditas. 1949;35S:446 50. Borntraeger; 1914. 106. Wiener AS. Heredity of the Rh blood type: VII. Additional family studies with R z ry – 71. Turesson G. The genotypical response of the plant species to the habitat. special reference to the genes and . Hereditas. 1949;35S:500 19. – Hereditas. 1922;3:211–350. 107. Kemp T. The rise of human genetics. Hereditas. 1949;35S:44 8. 72. Stamhuis IH, Monsen A. Kristine Bonnevie, Tine Tammes and Elizabeth 108. Hofsten N v. Ärftlighetslära. Föreläsningar vid Uppsala universitet 1916-1917. Schiemann in early genetics: emerging chances for a university career for Stockholm: Norstedts; 1919. (In Swedish). women. J Hist Biol. 2007;40:427–66. 109. Hofsten N v. The genetic effect of negative selection in man. Hereditas. – 73. Bonnevie K. Zur Analyse der Vererbungsfaktoren der Papillarmuster. 1951;37:157 265. Hereditas. 1923;4:221–30. 110. Norton HTJ, Appendix I. Punnett RC. Mimicry in butterflies. Cambridge: – 74. Bonnevie K. Papillarmuster und psychische Eigenschaften. Hereditas. 1927;9: Cambridge University Press; 1915. p. 154 6. Galeopsis tetrahit 180–92. 111. Müntzing A. Cytogenetic investigations on synthetic . – 75. Bonnevie K. Papillarmuster bei Linkshändigen. Hereditas 1933; 18:129-39. Hereditas. 1932;16:105 54. Berglund V. Sechs Fälle von hypotrichosis in einer Familie. Hereditas. 1924;5: 112. Westergaard M. Arvelighedslæren. Copenhagen: Munksgaard; 1953. (In 44–8. Danish). 113. Brainard J. Tainted names are removed from places of honor. Science. 2020; 76. Bonnier G. Heterozygosis in man as the supposed genetical basis of inclination to tuberculosis. Acta Zool. 1923;4:111–21. 369:122. 114. Tjio JH, Levan A. The chromosome number of man. Hereditas. 1956;42:1–6. 77. Bonnier G. De l’applicabilité de quelques formules statistiques à la biologie. 115. Caspersson T, Hultén M, Lindsten J, Zech L. Identification of chromosome Hereditas. 1927;10:237–40. bivalents in human male meiosis by quinacrine mustard florescence 78. Pettersson G, Bonnier G. Inherited sex-mosaic in man. Hereditas. 1937;22:49–69. analysis. Hereditas. 1971;67:147–9. 79. Bonnier G. Inherited sex-mosaic in man and the statistical treatment of such data. An answer to professor S. G Levit Hereditas. 1938;24:465–70. 80. Dahlberg G. Twins and heredity. Hereditas. 1923;4:27–32. Publisher’sNote 81. Dahlberg G. Eine neue Methode zur familienstatistischen Analyse bei der Springer Nature remains neutral with regard to jurisdictional claims in Vererbung. Hereditas. 1930;14:73–83. published maps and institutional affiliations. 82. Dahlberg G. Inzucht und Polyhybridtät beim Menschen. Hereditas. 1930b;14: 83–96. 83. Weinberg W. Zur Probandenmethode. Hereditas. 1931;15:120–6. 84. Dahlberg G, Diamant M. Inheritance of pneumatisation of the mastoid bone. Hereditas. 1945;31:441–56. 85. Berglund V. Sechs Fälle von Hypotrichosis in einer Familie. Hereditas. 1924;5: 44–8. 86. Ab MJ. Zur Erbanalyse der musikalischen Begabung. Hereditas. 1925;7:109–28. 87. Mjöen FA. Die Bedeutung der Tonhöhenunterschiedsempfindlichkeit für die Musikalität und ihr Verhalten bei der Vererbung. Hereditas. 1926;7:161–88. 88. Ribbing L. Sur la persistance d’un type crȃnien depuis l’ȃge de pierre jusqu’à nos jours dans une contrée suédoise. Hereditas. 1926;7:145–50. 89. Wahlund S. Zusammensetzung von Populationen und Korrelationserscheinungen vom Standpunkt der Vererbungslehre aus betrachtet. Hereditas. 1928;11:65–106.